Botanical Journal of the Linnean Society

SAUQUET, HERVÉ; DOYLE, JAMES A.; SCHARASCHKIN, TANYA; BORSCH, THOMAS; HILU, KHIDIR W.; CHATROU, LARS W.; LE THOMAS, ANNICK

doi: 10.1046/j.1095-8339.2003.00171.xpmid: N/A

AbstractMagnoliales, consisting of six families of tropical to warm-temperate woody angiosperms, were long considered the most archaic order of flowering plants, but molecular analyses nest them among other eumagnoliids. Based on separate and combined analyses of a morphological matrix (115 characters) and multiple molecular data sets (seven variable chloroplast loci and five more conserved genes; 14 536 aligned nucleotides), phylogenetic relationships were investigated simultaneously within Magnoliales and Myristicaceae, using Laurales, Winterales, and Piperales as outgroups. Despite apparent conflicts among data sets, parsimony and maximum likelihood analyses of combined data converged towards a fully resolved and well-supported topology, consistent with higher-level molecular analyses except for the position of Magnoliaceae: Myristicaceae + (Magnoliaceae + ((Degeneria+Galbulimima) + (Eupomatia+ Annonaceae))). Based on these results, we discuss morphological evolution in Magnoliales and show that several supposedly plesiomorphic traits are synapomorphies of Magnoliineae, the sister group of Myristicaceae (e.g. laminar stamens). Relationships within Annonaceae are also resolved with strong support (Anaxagorea basal, then ambavioids). In contrast, resolution of relationships within Myristicaceae is difficult and still incomplete, due to a very low level of molecular divergence within the family and a long stem lineage. However, our data provide good evidence that Mauloutchia is nested among other Afro-Malagasy genera, contradicting the view that its androecium and pollen are plesiomorphic

TAPLIN, JAMES R. D.; LOVETT, JON C.

doi: 10.1046/j.1095-8339.2003.00164.xpmid: N/A

AbstractIn order to investigate continental-scale patterns of plant species richness and rarity, distribution maps of 3661 plant species were digitized into a one degree grid of sub-Saharan Africa using the WORLDMAP computer programme. Cells with high species richness were also likely to be those containing the greatest number of species of restricted range, but areas such as the South African Cape and the Eastern Arc mountains were found to have more restricted-range species than predicted from their richness scores. The two environmental predictors which had the strongest individual relationships with both species richness and range-size rarity were absolute maximum annual temperature and mean monthly potential evapotranspiration. However, correlative predictive powers of these variables were low, with R=−0.58 and R=−0.54, respectively (P < 0.01). Multiple regression also failed to produce a strong explanatory model for observed continental-scale patterns of diversity. Spatial variability analysis showed that this was likely to be because different environmental parameters predicted different centres of richness and rarity. West African species richness was better predicted by absolute maximum annual temperature, whereas East African species richness was better predicted by mean monthly potential evapotranspiration.

PŠENIČKA, JOSEF; BEK, JIŘÍ; ZODROW, ERWIN L.; CLEAL, CHRISTOPHER J.; HEMSLEY, ALAN R.

doi: 10.1046/j.1095-8339.2003.00169.xpmid: N/A

AbstractSydneia manleyi gen. et sp. nov. is based on part of a fertile frond from the upper Westphalian D of the Sydney Coalfield, Nova Scotia, Canada. It has small synangia composed of laterally fused sporangia that are elongate and with a circular cross-section. The sporangia yielded variably sized monolete and trilete spores with laevigate and microspinate ornamentation; intermediate forms were also observed. The spores can be correlated with the sporae dispersae species Latosporites minutus, Punctatosporites oculus and Laevigatosporites minimus. Size distribution of the spores is variable and highly skewed, suggesting heterogeneity of the spores within the sporangium. Spore ultrastructure indicates that the fossil is part of a fern, and the morphology of the spores and synangia indicate marattialean affinities.

CHIAPELLA,, JORGE;PROBATOVA, NINA, S.

doi: 10.1046/j.1095-8339.2003.00167.xpmid: N/A

Abstract The Deschampsia cespitosa complex in Russia is revised. Related taxa common to Russia and adjacent regions (i.e. northern Europe, Arctic America and Japan) are also considered. Deschampsia cespitosa s.l. is characterized by open to less commonly rather contracted panicles, spikelets (1) 2 (3)-flowered, 4-toothed lemmas with the lateral teeth larger, and awns straight or bent, inserted from the base to the top of the lemmas. Due to extensive overlapping in diagnostic characters, it is not possible to accept the specific status for D. borealis (Traut.) Roshev., D. brevifolia R.Br., D. macrothyrsa Tatew. et Ohwi and D. obensis Roshev; instead, they are treated as subspecies of D. cespitosa (L.) Beauv. Morphological differences are insufficient for a clear differentiation among subspecies, and geographical distributions also need to be considered. A total of 14 subspecies is accepted. Observations on ecology, problems of specific and subspecific delimitation, a distribution map, and considerations of evolution of the complex are provided. One new combination is made, Deschampsia cespitosa ssp. sukatschewii (Popl.) Chiapella & Prob. geographical distribution, morphological variability, subspecies INTRODUCTION Deschampsia P. Beauv. (Poaceae: Aveneae) occurs mostly in wetlands, meadows and mountain grasslands of temperate and cold temperate regions, while in tropical areas it is restricted to high mountain regions. Since its description by Palisot de Beauvois (1812), the high morphological diversity of Deschampsia, particularly of D. cespitosa, and the lack of clear boundaries with related genera has resulted in much confusion. Several species now accepted as belonging to Deschampsia were placed under other genera: Periballia (Trinius, 1820), Campella (Link, 1827), Podinapus (Dulac, 1867), Avenella (Parlatore, 1850), Czerniaevia (Ledebour, 1853), Airidium (Steudel, 1855), Lerchenfeldia (Schur, 1866). The unclear limits in Deschampsia are not restricted to the generic level, but are also common in the infraspecific level differentiation of D. cespitosa. Several slightly different morphological variants of this species were treated as species (Polunin, 1959; Hultén, 1962; Kawano, 1963, 1966; Chrtek & Jirásek, 1965; Hess, Landolt & Hirzel, 1967; Clarke, 1978; Lid, 1985; Conert, 1987; Schmeil, 1988) but are today considered either as accepted synonyms of D. cespitosa (L.) P. Beauv. or as subspecies of D. cespitosa. After Lawrence (1945), Polunin (1959), Hultén (1962) and Kawano (1963, 1966), some taxa of uncertain status were treated as subspecies or varieties of the D. cespitosa‘complex’. In particular in Russia this approach was followed by Tzvelev, 1974), who reduced to subspecies of D. cespitosa four taxa previously considered species (D. alpina (L.) Roem. et Schult., D. brevifolia R. Br., D. mezensis Senjan.-Korcz et Korcz. and D. obensis Roshev.). In 1964 Tzvelev had given specific status to D. alpina (L.) Roem. et Schult., D. anadyrensis A.V. Vassil., D. brevifolia R. Br., D. borealis (Traut.) Roshev., D. glauca C. Hartm., D. obensis Roshev. and D. sukatschewii (Popl.) Roshev., estimating the number of species of the genus to be about 100. In the 1995 translation of his work, Tzvelev noted that (p. 150) ‘… almost all the arctic species of subgenus Deschampsia are very closely related and readily hybridize with one another, apparently producing fertile hybrids . . .’ and ‘. . . the complete lack of absolutely constant characters and insignificance of the differences between species possibly justifies the position taken by many authors right up to the present time that many species of subgenus Deschampsia are units of lower taxonomic rank – subspecies or even varieties’. In line with this, in 1976 Tzvelev reduced to subspecific status D. alpicola Rydb., D. koelerioides Regel, D. macrothyrsa (Tatew. et Ohwi) Kawano, D. pamirica Roshev., D. paramushirensis Honda, D. turczaninowii (Litv.) Spiosk. and D. wilhelmsii, thus reckoning only two species (D. cespitosa (L.) P. Beauv. and D. media (Gouan.) Roem. et Schult.) in the former Soviet Union and 20 species for all the genus (Tzvelev, 1976: 412). This tendency to lump related taxa into the D. cespitosa‘complex’ was not always followed in central Europe, where D. wibeliana (Sond.) Parl. and D. littoralis (Gaudin.) Reut. were considered separate species of the D. cespitosa (L.) P. Beauv. complex by Hess et al. (1967), Conert (1987) and Lauber & Wagner (1998), although Clarke (1978, 1980) had already reduced D. wibeliana (Sond.) Parl. to a subspecies. In northern Europe, Lid (1985) treated D. bottnica (Whalenb.) Trin. as separate from D. cespitosa (L.) P. Beauv. Probatova (1985) accepted ten taxa for the Russian Far East (D. cespitosa (L.) P. Beauv., D. beringensis Hultén, D. macrothyrsa (Tatew. et Ohwi) Kawano, D. obensis Roshev., D. tzvelevii, D. paramushirensis Honda, D. borealis (Traut.) Roshev., D. brevifolia R. Br., D. glauca C. Hartm., D. sukatschewii (Popl.) Roshev.); these were granted specific status on the basis of different ecological preferences and geographical areas. Czerepanov (1995) produced a check-list recognizing the specific status of D. alpina (L.) Roem. et Schult., D. beringensis Hultén, D. borealis (Traut.) Roshev., D. brevifolia R. Br., D. glauca C. Hartm., D. koelerioides Regel, D. macrothyrsa (Tatew. et Ohwi) Kawano, D. mezensis Senjan.-Korcz et Korcz., D. obensis Roshev., D. pamirica, D. paramushirensis Honda, D. sukatschewii (Popl.) Roshev., D. turczaninowi (Litv.) Spiosk., D. tzvelevii Prob. and D. vodopjanoviae O.D. Nikif. Recently, Chiapella (2000a) analysed the morphological variability of Deschampsia cespitosa for central and northern Europe and recognized a total of six subspecies of which D. cespitosa ssp. alpina (L.) Tzvelev, D. cespitosa ssp. glauca (C. Hartm.) C. Hartm. and D. cespitosa ssp. parviflora (Thuill.) K. Richt. are found in the European part of Russia. The objective of the present work is to provide an update of the taxa included in Deschampsia cespitosa s.l. present in Russia and adjacent regions, including observations on taxon delimitation, nomenclature problems, ecology and geographical distribution. MATERIAL AND METHODS Classical taxonomic methods were used. Material examined belongs to B, BM, BP, GZU, K, LE, LEU, M, MHA, P, S, US, VLA, W and WU (Holmgren, Holmgren & Hirzel, 1990). Delimitation of taxa in Deschampsia cespitosa presented practical problems, because of the likely existence of hybridization (Kawano, 1966; Clarke, 1980; Hedberg, 1986) and different ploidy levels within the same phenotype make taxon boundaries indistinct (Barlow-Irick, 1997). The existence of abundant transitional forms between varieties suggested avoiding the use of this category, due to the impossibility of defining clear boundaries between the taxa. The only infraspecific category recognized was the subspecies, defined following Du Rietz (1930): a subspecies was defined as the regional facies of a species correlated with a restricted geographical distribution (allopatric). RESULTS Geographic distribution The geographical distribution of all the taxa except D. cespitosa ssp. cespitosa (L.) P. Beauv., D. cespitosa ssp. parviflora (Thuill.) K. Richt. and D. cespitosa ssp. sukatschewii (Popl.) Chiapella et Prob. was determined (Fig. 1) on the basis of herbarium material and published sources (Hultén, 1927, 1937, 1941, 1962; Kawano, 1963; Tzvelev, 1974, 1976, 1995; Probatova, 1985). Subspecies cespitosa and parviflora are widespread in Europe and in the European part of Russia, while the distribution of ssp. sukatschewii is not entirely clear yet; most of the other subspecies have allopatric distributions with zones of sympatry. Figure 1. Open in new tabDownload slide Distribution of Deschampsia cespitosa ssp. alpina, D. cespitosa ssp. beringensis, D. cespitosa ssp. borealis, D. cespitosa ssp. brevifolia, D. cespitosa ssp. glauca, D. cespitosa ssp. macrothyrsa, D. cespitosa ssp. mezensis, D. cespitosa ssp. obensis, D. cespitosa ssp. orientalis, D. cespitosa ssp. submutica and D. cespitosa ssp. turczaninowii. Figure 1. Open in new tabDownload slide Distribution of Deschampsia cespitosa ssp. alpina, D. cespitosa ssp. beringensis, D. cespitosa ssp. borealis, D. cespitosa ssp. brevifolia, D. cespitosa ssp. glauca, D. cespitosa ssp. macrothyrsa, D. cespitosa ssp. mezensis, D. cespitosa ssp. obensis, D. cespitosa ssp. orientalis, D. cespitosa ssp. submutica and D. cespitosa ssp. turczaninowii. Taxonomy Deschampsia cespitosa (L.) P. Beauv., Ess. Agrostogr.: 91. 1812 Description: Perennial densely tufted grass with extremely variable habit; culms erect, 5–150 cm high; leafy at the base, sheaths glabrous with margins membranous to scarious. ligules obtuse to acute, 2–12 mm long, membraneous or scarious. leaves elongate, linear, flat or conduplicate, 2–34 cm long, 1–5 mm wide, adaxially scabrous, abaxially glabrous, with 6–8 visible ribs, rough to the touch, margins often scarious. panicles wide, open, frequently nodding, 3–37 cm long, 1.5–7 cm wide, rarely contracted, peduncles glabrous or minutely scabrous. spikelets (1)−2–(3)-flowered, compressed, purple with green and/or gold (or a mix); rachilla usually with abundant hairs, callus hairs short, about 1/3 the length of the lemma; glumes usually covering the florets; lower glume lanceolate, 2–7 mm long, membraneous to scarious, acute, entire, 1–3 nerved; upper glume narrowly elliptic, 2–6 mm long, acute, 3-nerved, with midnerve glabrous or scaberulous; lemma narrowly oblong, 2.5–3.5 mm long (3)−4–(5) toothed to erose-toothed, with lateral teeth larger, most rarely all equal or the central larger (1)-3–5 nerved, membraneous; awn straight or bent, weak, commonly not twisted, 1–8 mm long, inserted usually at the basal or median portions of the lemma, scabrous; palea hyaline, 2–3 mm long, 2-keeled, with scabrous keels. anthers 1–2 mm long, yellow or purple. caryopsis 0.5–1 mm long, ovoid, brownish. 1. Deschampsia cespitosa ssp. alpina (L.) Tzvelev, Fl. Sev.-Vost. Europ. 1: 141. 1974. Aira alpina L., Sp. Pl. 65. 1753. Deschampsia alpina (L.) Roem. et Schult., Syst. Veg. 2: 686. 1817. Aira major ssp. alpina (L.) Syme ex Sowerby Engl. Bot., ed. 3, 11: 65. 1877. Aira caespitosa ssp. alpina (L.) Hook. f. Student. fl. Brit. Isl. 3: 437. 1870. type: Sweden, Torne Lappmark, Mt. Nuolja, c. 700–800 m, N.D. Simpson 50133, 25.vii.1950 (Lectotype BM! designated by Cope in Cafferty et al. Taxon 49: 243 (2000)), not Gaudin 1828. Aira laevigata Sm., Trans. Linn. Soc. 10: 334. 1810. Deschampsia laevigata (Sm.) Sm ex Roem. et Schult., Syst. Veg. 2: 687. 1817. type: Scotland, High mountains of Clova, Angusshire, and the coast near Dundee. Aira cespitosa var. alpina Gaudin, Fl. helv. 1: 323. 1828. Deschampsia cespitosa var. alpina Gaudin, Taschenb. Schweiz. Bot. 1869. non L. 1753. Deschampsia cespitosa var. alpina Schur, Oesterr. Bot. Z. 9: 326. 1859. type: Switzerland, auf Alpentriften längs der ganzen Kette, wo dieses Gras auf den Glimmerschiefergebirgen die Hauptvegetation bildet, von 5000 bi 7000′ Elevation, Hochmoorboden, non Aira alpina L. 1753. Deschampsia cespitosa var. alpina Honda, J. Fac. Sci. Tokyo 3: 139. 1930. non L. 1753. Description: Plants 8–40-(65) cm; ligules 2–6 mm long; blades 2–5 (7) cm long, 0.5–2 mm wide. Panicles rather contracted, 5–15–(38) cm long; spikelets 2–5.5 mm long (1)−2–3 flowered; lower glume 2–5 mm long; upper glume 3–5.5 mm long; lemmas 2–5 mm long; awns straight, not twisted, 1–3 mm long, inserted in the upper or medium third of the lemma. Chromosome number: 2n = 26, 35–38, 38–39, 39, 41, 48, 49, 49 + 2B, 49–52, 50, 52, 56 (Fedorov, 1969; Goldblatt, 1984; Elven, 1994). Distribution: Russia: Kola Peninsula and Novaya Zemlya Island; mountains of Scandinavia. Key to the subspecies ofDeschampsia cespitosapresent in Russia, modified afterProbatova (1985),andChiapella (2000b) 1. Spikelets 1–2-flowered; Central and Eastern Europe, Russia, European part (introduced in the Russian Far East) 2 1′. Spikelets 2–3 flowered; Siberia, Russian Far East 3 2. Spikelets 3–4.5 mm; awns not exceeding the top of their lemmas; panicles with fewer than 100 spikelets, green, dark green or violaceous, or a mix D. cespitosa ssp. cespitosa (s.s.) 2′. Spikelets 2–3.5 mm; awns exceeding or not the top of the their lemmas; panicles with 100–150 pale green spikelets D. cespitosa ssp. parviflora 3. Panicles rather contracted 4 3′. Panicles more open and lax 11 4. Spikelets often pseudoviviparous; mountains of Scandinavia, Novaya Zemlya Island. D. cespitosa ssp. alpina 4′. Spikelets never (or rarely) pseudoviviparous 5 5. Awns very short, weak, c. 1 mm long; Russia, Siberia, Olenek river D. cespitosa ssp. submutica 5′. Awns more than 2 mm long 6 6. Awns 8–12 mm long; Alaska, Kamchatka peninsula, Commander Islands, Kurile Islands D. cespitosa ssp. beringensis 6′. Awns 2–8 mm long 7 7. Awns inserted in the lower third of the lemma or at its base 8 7′. Awns inserted in the middle of the lemma 9 8. Ligules 6–9 mm long; Eastern Siberia (Baikal Lake) D. cespitosa ssp. turczaninowii 8′. Ligules 1.5–4.5 mm long 10 9. Blades 5.5–7 cm long; ligules 3–4 mm long; Arctic Russia, Western and Eastern Siberia, Novaya Zemlya Island, Verkhoturova and Karaginsky Islands D. cespitosa ssp. borealis 9′. Blades 10–20 cm long; ligules 4–5 mm long; Russia (European part), along Mezen and Vashka rivers D. cespitosa ssp. mezensis 10. Awns 4–6 mm long; spikelets 2–3 flowered; arctic North America, Greenland, Russia (Chukotka peninsula) D. cespitosa ssp. brevifolia 10′. Awns 1.5–3.5 mm long; spikelets 2-flowered; northern Sweden, Finland, Russia (European part) D. cespitosa ssp. glauca 11. Awns bent; blades 5–8 cm long; sand dunes of the Arctic Sea, estuaries of the Obj and Yenissei rivers D. cespitosa ssp. obensis 11′. Awns straight; blades 5–25 cm long 12 12. Ligules 4–5 mm; plants with rhizomes; marshy sea coastal meadows in Japan (Hokkaido), and Russia (The sea of Okhotsk, Sakhalin and the Kuriles) D. cespitosa ssp. macrothyrsa 12′. Ligules 2–3 mm long; plants without rhizomes 13 13. Branches of the panicle glabrous; spikelets 2-flowered, 2.5–3.5 (4) mm; Russia (Eastern Siberia and continental part of the Far East) D. cespitosa ssp. sukatschewii 13′. Branches of the panicle scabrous; spikelets 2-(3)-flowered 3–5 mm long; Russia, Kamchatka Peninsula, Yakutia; Japan, Hokkaido, Honshu, Kiusiu D. cespitosa ssp. orientalis Ecology: Mountain stony slopes in Scandinavia, tundra in Kola and Novay Zemlya. Specimens examined:russia. Novaja Zemlja, Insula Septentrionalis, in litore arenoso-schistoso prope Matotschkin, A. Tolmatschev 4779, 2.ix.1927 (K); idem, Matotschkin-Schar, L. Limanowski 72, vii.1909 (LE); Kola Peninsula, Murmansk Region, Lovozersky Mountains, Mt Karnasurta, H. Semenova 179, 13.viii.1959 (LE). finland. Petsamo, Petsamontunturit, A. Cajander, 19.viii.1929 (S); Lapponia Enontekiensis, pratis alpinis Montis Raasa, I. Montell, 5.viii.1912 (S). norway. Dovre, Kongsvold, P. Olsson, 18.viii.1887 (GZU); Tromsö, A. Noto 226, 23.i.1901 (GZU); Nordfjord, Loenbran, M. Sonden s.n. 7.vii.1899 (S); Spitzbergen, Kongsfjord, H. Hertel 16691, 22.vii.1975 (M); Recherche Bay, G. Andersson et H. Hesselman 246, 13.vii.1898 (S); Stans Foreland, Krauss Haffen, A. Birula 18, 17.viii.1899 (LEU); Finnmark, Hammerfest, H. Merxmüller et W. Wiedmann 258/59, 18.viii.1959 (M); Alpibus Norvegicus, Herb. Trinius 1875–11 (LE). sweden. Härjedalen, Tännäs, Funäsdalen, M. Östman, 21.viii.1910 (S); Jämtland, Undersäker, Sylfjällen, T. Arwidsson s.n., 15.vii.1943 (S); Äsele Lappmark, Daunatjäkko, Vilhemina socken, S. Nordenstam, 28.vii.1939 (S); Lycksele Lappmark, Tärna socken, E. Asplund, 24.vii.1937 (S); Torne Lappmark, Kiruna, A. Stork, 12.vii.1960 (S). Observation: The spikelets are often deformed due to pseudovivipary, a common feature in Scandinavian mountain populations but with no clear relation to either environment or genetic background (Hedberg, 1986). This subspecies has the smallest panicles within the D. cespitosa group. 2. Deschampsia cespitosa ssp. beringensis (Hultén) W. E. Lawr., Amer. J. Bot. 32: 302. 1945. D. beringensis Hultén, Kungl. Svenska Vetenskapsakad. Handl. 3: 107. 1927. type: Russia, Bering Island, Sandberg, 18.viii.1921 (holotype S!). Deschampsia cespitosa var. arctica Vasey, USDA Division Bot. Bull. 13: 45. 1892. type: USA, Coast of Alaska, Saint Paul Island, Merriam (holotype US?, n.v). Deschampsia aleutica Tatew. et Ohwi, J. Fac. Agr. Sapporo University 36: 95. 1934. type: USA, Aleutian Islands. Atka, hab. Nazan, M. Tatewaki et K. Takahashi 14485, 1927 (holotype HAK?, n.v). Deschampsia beringensis var. atkensis Hultén, Fl. Aleut. is. 81. 1937. type: USA, Aleutian Islands, Atka, E. Hultén 6562, 12.vii.1932 (holotype S!). Description: Plants 25–80 cm high, stems usually thick; ligules acute, 8–12 mm long; blades flat, 5–12 cm long, 2–4 mm wide, minutely scabrous. Panicles rather contracted, 9–20 cm long; spikelets 4–7 mm long, 2–3-flowered; glumes narrowly lanceolate, equal, 4–7 mm long, 3-nerved; lemma 3–4.5 mm long; awn straight, 3–6 mm long, inserted in or near the base of the lemma. Chromosome number: 2n = 26, 42 (Sokolovskaya & Probatova, 1975). Both counts were made on plants from Bering Island, although the count 2n = 42 probably does not belong to this taxon. Distribution: Russia: southern part of Kamchatka Peninsula, Commander Islands, the Kuriles. USA: Aleutian Islands, Alaska. Canada: west coast. Ecology: Dry meadows in tundras, coastal sand dunes, slopes near the sea. Specimens examined:russia. Kurile Islands, Urup Island, Kastricum Peninsula, meadow, Meljnikova et Butovsky 326, 1968 (MHA); Commander Islands, Island Medny, near Preobrazhenskaya Village, dry meadow, Kardakov, 27.viii.1930 (LE); Kamchatka, s. colec., 1879 (LE). usa. Alaska, Aleutian Islands, Unimak Island, False Pass, W.J. Eyerdam 2130, 5.viii.1932 (US); Gulf of Alaska, Middleton Island, upper edge of beach below bluff, above normal high tide line at base of small bluff, J.H. Thomas 6309 A, 2.viii.1956 (US); Prince William Sd., Hinchinbrook, I.L. Norberg, US 2238673 (US). canada. British Columbia, Queen Charlotte Island, west coast of Moresby Island, common on rocky grassy slopes from 400 feet well above treeline, J.A. Calder et R.L. Taylor 23605, 16.viii.1957 (B), Graham Island, Port Clements, common or sedge flats along margin of creek, J.A. Calder et al. 22108, 3.vii.1957 (B). Observation 1:Hultén (1927) regarded this taxon as close to D. cespitosa ssp. bottnica (Wahlenb.) Tzvelev, which is found on the Swedish and Finnish coasts of the Gulf of Bothnia. Besides the disjunct geographical distribution, D. cespitosa ssp. beringensis (Hultén) W. E. Lawr. differs from ssp. bottnica in having smaller plants, 2–(3)-flowered spikelets, shorter flat blades and a densely pilose callus, with hairs often reaching half way up the lemma. Observation 2: In 1937 Hultén described D. beringensis var. atkensis Hultén on the basis of a specimen with awns inserted in the middle of the lemmas. This character, however, is not constant, as the type specimen preserved in S (Aleutian Islands, Atka, E. Hultén 6562, 12.vii.1932) shows also lemmas with awns inserted in the basal third. 3. Deschampsia cespitosa ssp. borealis (Traut.) Á. Löve et D. Löve, Op. Bot. 5: 65. 1961. Aira cespitosa var. borealis Traut., Act. Hort. Petrop. 1: 86. 1871. Deschampsia borealis (Traut.) Roshev. in Tolmatch., Fl. Tsentr. Taimyra 8: 93. 1932. type: Russia, Ad fluvius Tajmyr, Middendorff, 1843 (holotype LE!). Description: Plants 5–30 cm high, often forming dense tufts with numerous old sheaths; ligules acute, 3–4 mm long, partially hyaline; blades flat to rather folded, 5–7 cm long, 0.5–1.5 mm wide, scabrous along the nerves at the adaxial side. Panicles contracted, 3–6 cm long; spikelets 3–4 mm long, 2(−3)-flowered; glumes roughly equal, 3–4 mm long, serrate near the tip; lemma 2–3 mm, hyaline at the distal portion; awn straight, 2–3 mm long, inserted in or slightly below the middle of the lemma, hairs of the callus often reaching 1/3 way up the lemma. Chromosome number: 2n = 24–26, 26, 28, 42, 52 (Zhukova & Petrovsky, 1972; Goldblatt, 1981, 1984; Takhtadzjan, 1993). Distribution: Russia: Novaya Zemlya Island, Arctic Siberia and Chukotka Peninsula, Bering Sea islands (Verkhoturova and Karaginsky Islands). Ecology: Gravelly shores of rivulets and small rivers, moss tundras and swampy meadows. Specimens examined:russia. Novaya Zemlja, insula Septentrionalis, sinus Russkaja Gavanj, V. Savicz, 8.vii.1931 (LE); Yamal Peninsula, river Matjuajako, meadow stream, affluent of river, O. Rebristaja, 5.viii.1983 (LE); Arctic Siberia, Dicksons hamn, 73°28′N, 80°58′W, F.R. Kjelman, 6–10.viii.1878 (K); Tajmyr Peninsula, east part of the peninsula, Maria Pronschischeva Bay, northern coast of Vetvisty rivulet in its lower part, N.V. Matvejeva 558, 12.viii.1972 (MHA); Tajmyr Peninsula, north coast of Tajmyrskoje Lake, Yamu-Baikura, wet tundra with Dryas and Carex spp., M.I. Vellikainen s.n., 13.viii.1947 (MHA); Yakutia, Begichev Island, N.V. Matvejeva 733, 22.viii.1972 (BM); Yakutia, lower part of Lena River, in vicinity of Tiksi Bay, stony place along a small river, A.P. Sokolovskaya 129, 1.viii.1948 (VLA); Chukotka, Chaunski region, K. Yakowlew, 12.ix.1937 (LE); Magadan Province, regio Tschukotsky, distr. Iultinsky, Polarny, tundra fruticuloso-herbida maculata, sola argillosa, copiose, V.V. Petrovsky 5604, 13.viii.1966 (B, BM, K, US); Kamchatskaya Region, Oljutorsky district, Verkhoturova Island, S.S. Kharkevich et T.G. Buch, 26.vii. 1975 and 27.vii. 1975 (VLA); Kamchatskaya Region, Karaginsky district, Karaginsky Island, W slope of mountain Vysokaya, tussock swamp meadow, common, S.S. Kharchevich et T.G. Buch 1043b, 22.viii.1976 (VLA); Herb. Institute Bot. Acad. Sc. URSS, 13.viii. 1948 US 2524228 (US); A.A. Korobkov, 4.viii.1975 (B). Observation 1: very similar to D. cespitosa ssp. brevifolia (R.Br.) Tzvelev. ssp. borealis (Traut.) Á. Löve et D. Löve is widely distributed in the Russian Arctic and it differs from ssp. brevifolia in having smaller plants and panicles and shorter leaves. Ssp. borealis (Traut.) Á. Löve et D. Löve was described from a plant of the Tajmyr River in the Tajmyr Peninsula, while ssp. brevifolia was described from the Canadian Arctic (Melville Island). In this work the epithet borealis is reserved for smaller plants occurring in the Russian Arctic, while brevifolia is used for plants of Greenland, Arctic America and Chukotka Peninsula. Observation 2: ssp. brevifolia and ssp. borealis (Traut.) Á. Löve et D. Löve are tentatively kept as separate subspecies, although there is a transition gradient between them. Tzvelev (1964, 1995) gave to both taxa the rank of species, but the existence of intermediates and the geographical distribution suggest that the rank of subspecies is more suitable. 4. Deschampsia cespitosa ssp. brevifolia (R. Br.) Tzvelev, Fl. Sev.-Vost. Evrop. Chasti SSSR 1: 141. 1974. Deschampsia brevifolia R. Br., Suppl. App. Parry's Voy. 191. 1821. Aira arctica Spreng., Syst. Veg. 4: 32. 1827. Aira cespitosa var. arctica (Spreng.) Thunb. ex A. Gray, Proc. Acad. Nat. Sci. Philadelphia 1863: 78. 1863. Deschampsia arctica (Spreng.) Merr., Rhodora 4: 143. 1902. D. arctica (Spreng.) Schischkin, Fl. Sib. Occid. 232. 1928. type: Canada ‘Arctic America’, Melville Island, Fischer s.n. (holotype Herb. Trinius LE! isotypes BM! S!). Description: Plants 9–55 cm high forming a sward; ligules acute, 1.5–4 mm long, hyaline; blades 2–10 cm long, 0.5–1.5 mm wide, convolute, rigid, acute, most rarely flat. Panicles rather contracted, 2.5–15 cm long; spikelets 3–6 mm long, 2–3-flowered; glumes roughly equal, 3–6 mm long; lemma 4–6 mm long; awn straight 2–4 mm long, inserted in the medial and more rarely in the basal third of the lemma. Chromosome number: 2n = 26, 27, 28, c. 50, 52 (Bowden, 1960; Fedorov, 1969; Zhukova, Petrovsky & Plieva, 1973; Petrovsky & Zhukova, 1981; Goldblatt, 1988; Takhtadzjan, 1993). Distribution: Russia: Chukotka Peninsula, Wrangel Island (?). USA: Alaska. Canada: Yukon, Arctic Canada. Greenland. Ecology: Tundra on rocky and sandy soils. Specimens examined:russia. Chukotka, Beringovsky district, middle part of Vaamochka River, 4.5 km NNE from the mouth of Vaapveyem River, small places with gravel and clay on the terrace, C.A. Balandin et al. 07/02, 28.vii.1985 (VLA). usa. Alaska, Prudhoe Bay area, NE of Deadhorse Airport, 70°18′N 148°20′W, W.R. Marris 19943, 31.vii.1980 (BM); Alaska, Seward Peninsula, Cape Prince of Wales, 65°48′N, 168°W, sea level to 700 msm, dunes, T. Kelso et al. 106, 24.vi.1979 (VLA). canada. Baffin island, Clyde River, J.L. Hanham 237a, 26.viii.1934 (BM); North-West Territories, Ellesmere Island, Hazen camp, 81°49′N, 71°21′W, densely caespitose, occasional, only on intermittently wet muddy delta, D.B.O. Savile 4827, 6.viii.1962 (US); Chesterfield, West Coast of Hudson Bay, A.E. Porsild 6133, 9.ix.1930 (S); Victoria Island, Cambridge Bay, W of Wes End, side of wet gulley, R. Marris 14743, 19.viii.1977 (BM); Yukon Territories, British Mts., 69°03′N 140°24′W, 2500 ft. broken rock and turf, W.J. Cody 27428, 9.viii.1980 (BM). greenland. Scoresby Island, Fleming fjord, Eddenfugle Dal, 71°53′N 22°40′W, R. Marris s.n., 18.viii.1963 (BM); Heilprind Land, Brønlund Fjord, 82°10′N 31°W, moist sandy soil in river delta, K. Holmen 8212, 10.viii.1950 (BM); Geographic Society Island, hab. in ripa arenosa, 72°44′N 22°30′W, 200 m, E. Einarsson et K. Holmen 12, chromosome number on label 2n = 26, 27.viii.1958 (BM); Thule, Firz Clavence, R. Platt et A.J. Boucot 188, 5.viii.1947 (US); Peary Land, Nedre Midsommersø, 82°13′N, 33°40′W, B. Fredskild 253 25.vii.1963 (US); Disko, Fortune Bay, S. Laegaard 328, 10.viii.1962 (S); Kangerdluarssuk, M. Möller 707, 28.vii.1981 (S). Observation 1: The chromosome count 2n = 52 (Petrovsky & Zhukova, 1981) was made on plants of Wrangel Island, which is here considered marginal for the distribution of this taxon. Tzvelev (1995) considered this taxon as also occurring in the Russian Arctic, but the specimens observed at LE are closer to ssp. borealis (Traut.) Á. Löve et D. Löve (see Observation in ssp. borealis). Some specimens with rigid short leaves typical of ssp. brevifolia have open panicles as in typical D. cespitosa. Observation 2:Aiken, Consaul & Dallwitz (2000) mentioned this taxon as an indicator of disturbed places, either by natural causes (floods, erosion) or human influence. The ability to colonize disturbed habitats has so far been reported only for tetraploid forms of D. cespitosa ssp. alpina in Great Britain (Rothera & Davy, 1986) and D. mackenziena in sand dunes in Canada (Raup & Argus, 1982; Purdy & Bayer, 1995). This is also the case of diploid ssp. cespitosa in the Russian Far East. 5. Deschampsia cespitosa (L.) P. Beauv. ssp. cespitosa, Ess. Agrostogr. 91. 1812. Aira cespitosa L., Sp. Pl. 64. 1753. Agrostis caespitosa (L.) Salisb., Prodr. Stirp. Chap. Allerton. 25. 1796. Campella cespitosa (L.) Link, Hort. berol. 1: 122. 1827. Avena cespitosa (L.) Kuntze, Taschen-Fl. Leipzig: 45. 1867. Podionapus caespitosus (L.) Dulac, Fl. Hautes-Pyrénées: 82. 1867. Aira major ssp. cespitosa (L.) Syme ex Sowerby, Engl. Bot. ed. 3, 11: 64. 1877. type: Europa, in partis cultis et fertilibus (holotype LINN; microfiche S!). Deschampsia biebersteiniana Schult. Mant. 2: 380. 1824. Aira biebarsteiniana (Schult.) Steud. Syn. pl. glumac. 1: 219. 1854. type: Caucasus, Circa acidulam Nartsana (Herb. Trinius?, n.v.). Aira cespitosa var. genuina Rchb., Icon. 1: pl. 96. 1834. (nom. nud.). Deschampsia cespitosa var. genuina (Rchb.) O.H. Volk, Bot. Jahrb. Syst. 47: 312. 1912. D. cespitosa ssp. genuina (Rchb.) W.E. Lawr., Amer. J. Bot. 32: 302. 1945. Aira andraei Auersw., Bot. Zeitung (Leipzig) 14: 205. 1856. type: Czech Republic, Siebenburger Alpen von Fogarasch, auf Krystall. Schieferen um die Gresbäche unweit, Stina Zirna gegen 5600′. Deschampsia andraei (Auersw.) Schur., Enum. pl. Transsilv. 753. 1866. A. cespitosa var. andraei (Auersw.) Grecescu, Consp. fl. Roman. 611. 1898. type: Romania, auf Triften der Alpen, Moorboden, auf dem Retyezat der Hunyader Alpen; auf den Kalkalpen bei Kronstadt. Aira major Syme ex Sowerby, Engl. Bot. 11: 63. 1873. Deschampsia cespitosa ssp. gaudini K. Richt., Pl. eur. 1: 56. 1890. D. cespitosa var gaudini (K. Richt.) Husn., Graminées 2: 34. 1897. Deschampsia ambigua P. Beauv. ex B.D. Jacks., Index Kewensis 1: 735. 1893. type: Canada, Mistassins. Deschampsia cespitosa var. alpina Vasey ex Beal Grass. N. Amer. 2: 368. 1896. non D. cespitosa var. alpina Gaudin 1869; non D. alpina (L.) Roem. et Schult. 1817. Deschampsia anadyrensis A.V. Vassil., Bot. Mater. Gerb. Bot. Institute Acad. Nauk SSSR 8: 68. 1940. Deschampsia cespitosa ssp. anadyrensis (A.V. Vassil.) Á. Löve et D. Löve, Bot. Not. 128: 503. 1975. type: Russia, Anadyr basin, lower course of Anadyr, opposite Krasnoje Lake, A.V. Vassiljev 1591, 10.viii.1933 (holotype LE!). Deschampsia cespitosa ssp. alpicola Chrtek et V. Jirásek, Acta University Carol. Biol. 3: 207. 1965. D. cespitosa var. alpicola (Rydb.) D. Löve et B.M. Kapoor, Arct. Alp. Res. 3: 143. 1971. D. cespitosa f. alpicola (Chrtek et V. Jirásek) Soó, Acta Bot. Acad. Sci. Hung.17: 120. 1971. type: Czech Republic, Kepernik, Goldstein, A. Oborny, 20.viii.1873 (holotype PR, n.v.). Deschampsia austrobohemica Deyl, Fol. Geobot. et Phyto. (Praha) 19: 101. 1984. D. cespitosa ssp. austrobohemica (Deyl) Conert, Illus. Flora Mittel. 305. 1987. type: Czech Republic, Bohemia Australis, Distr. Blatná, in pratis prope vicum Drabenicky, M. Deyl, 27.vi.1972 (holotype PR!). Deschampsia vodopjanoviae O.D. Nikif. Bot. Zhur. 72 (12): 1666. 1987. type: Russia, Putorana Plateau, lacus Chaja-Kjuel, in regione subalpina, declive schistosum ad flumen, S. Andrulaitis et Z. Malyscheva, 9.viii.1972 (holotype LE!). Description: Plants (30)−40–80–(120) cm high, leafy at the base; ligules obtuse to acute, 2–8 mm long, membraneous or scarious; blades linear, flat or conduplicate, 2–25 cm long, 2–4–(5) mm wide, adaxially scabrous, abaxially glabrous, the adaxial side with evident veins. Panicles wide, open, 8–25 cm long, 4–10 cm wide, rarely contracted, broadly spread during anthesis; spikelets (1)−2–(3)-flowered; glumes slightly unequal, lower glume lanceolate, 2–4.5 mm long, 1–3 nerved; upper glume narrowly elliptic, 2–6 mm long, 3-nerved; lemma oblong, 2.5–3.5 mm long, toothed to erose-toothed; awn straight or bent, 1–8 mm long, inserted usually at the basal or median portions of the lemma. Chromosome number: 2n = 18, 24, 24–25, 26, 26 + 1–2B, 27, 28, c. 39, 52 (Albers, 1972, 1980; Fedorov, 1969; Goldblatt, 1981, 1984, 1985, 1988; Goldblatt & Johnson, 1990, 1994, 1996, 1998, 2000). Distribution: Subspecies widely distributed in almost all cold temperate areas of both hemispheres. Ecology: Wet meadows, bogs, along streams and water courses, and on lake shores. From sea level up to about 3000 m. Specimens examined:russia. Rossia Arctica, Lapponia tulomensis, litus Murmanicum prope Alexandrovsk, 22.viii.1927, P. Nikolsky 6155 (LE); Moskovskaya Region, Ruzsky district, in vicinity of Glubokoye Lake, E.E. Gogina et A.E. Matsenko, 29.vi.1980 (MHA); Siberia, Jenisei, M. Bremer, 1876 (S); Lake Baikal, Zungoi Island, H.H. Iltis et al. 317, 1.vii.1979 (S); Magadanskaya Region, 30 km from Magadan sity, Snezhnaya Dolina settlement, V.P. Seledets, 10.viii.1971; Kamchatka Peninsula, south-east part, in vicinity of Nalychevskije hot springs, Gorjachaya River, in water, partly submerged, V.V. Yakubov, 3.ix.1995; Sakhalin, Tymovsky district, in vicinity of Longari railway station, E.M. Egorova, 24.vii.1978 (MHA); Primorsky Territory, Chugujevsky district, nearby Novo-Mikhajlovka settlement, grazed lawn, V. Kuzjminykh, 26.vi.1969 (MHA); Primorsky Territory, Shkotovsky district, Anissimovka railway station, in the edge of the village on the way to Tigrovaya station, T.I. Nechajeva, 10.vii.1973 (MHA). usa. Alaska, Nome, open ground along road in hills, A. S. Hitchcock 1425, 29.x.1909 (P). estonia. Vormsi Island, Borlay, J. Pedaste 24, 8.vii.1975 (B). china. Yunnan Province, eastern slopes of L'akiang snow range, Yangtze watershed, J.F. Rock 10641, 1924 (P). Observation: A morphologically very variable taxon with different chromosome numbers reported throughout the world, although 2n = 26 seems to be the most common. 6. Deschampsia cespitosa ssp. glauca (C. Hartm.) C. Hartm., Sv. norsk exc.-fl. 15. 1846. Deschampsia glauca C. Hartm., Handb. Skand. fl. 448. 1820. Deschampsia cespitosa var. glauca (C. Hartm.) Sam., Sv. Bot. Tidskr. 13: 253. 1919. type: Sweden, Jemtland, Backman s.n. (UPS, n.v.). Description: Plants 30–50 cm high; ligules 3–4.5 mm long; blades 4–8.5 cm long, 1–2 mm wide, convolute or folded, rough, glaucous often pointed downwards. Panicles somewhat contracted, 10–14 (18) cm long; spikelets 2–4.5 mm long, 2-flowered; lower glume 2–3 mm long; upper glume 3–4.5 mm long; lemma 2–3 mm long; awns straight, not twisted, 1.5–3.5 mm long, inserted in the basal third of the lemma. Chromosome number: 2n = 26, 48–50, c. 52, 52 (Takhtadzjan, 1993; Elven, 1994). Distribution: Scandinavia (excluding Denmark). Russia: Arctic European and Asiatic (rare), Wrangel Island (?). Ecology: Tundra rocky soils, along rivulets and streams. Specimens examined:russia. Tajmyr Peninsula, east part of the peninsula, Byrranga Mountains, middle part of Njunjkarakutari River, near the lake Poimennoye, I.N. Pospelov 98–2, 5.viii.1998 (MHA); Tajmyr Peninsula, central part, Byrranga Mountains, Fadjukuda River, salty place near the Snezhny creek, I.N. Pospelov 97–266, 14.viii.1997 (MHA); Wrangel Island, W. Lewkowsky et Tikhmeniew, 15.vii.1972 (LE). finland. Lapponia, Ponoj, A.J. Malmberg, viii.1870 (S). sweden. Lapponia, Abisko, Torneträsk, A. Palmgren GZU 142396, 1910 (GZU); Lycksele Lappmark, Lycksele Socken, E. Asplund, 14.vii.1937 (S); Lule Lappmark, Kvikkjokks Kapellförs, S. Selander, 7.viii.1946 (S); Torne Lappmark, H. Dahlstedt, 5.viii.1910 (S); Jemtland, Lugnvik, Storsjön lake, G. Öhrstedt, 5.viii.1921 (S). Observation 1: Plants with 2n = 52 have been reported pseudoviviparous (Elven, 1994). The counts 2n = 52 by Zhukova & Petrovsky (1976) and Petrovsky & Zhukova (1981) were made on plants of Chukotka and Wrangel Island. Observation 2: Originally described from Emtland province in northern Sweden, it is found also in Finland. Tzvelev (1976) cited it also for Russia, where it becomes sparse to the east of the Urals, being gradually replaced by ssp. borealis (Traut.) Á. Löve et D. Löve. The Russian material studied at LE is closer to ssp. borealis than to ssp. glauca (C. Hartm.) C. Hartm.; however, the specimen Lewkowsky et Tikhmeniew (LE), collected in Wrangel Island agrees, well with European material of D. cespitosa ssp. glauca. Observation 3:Aiken et al. (2000) mentioned this taxon as ‘uncertain’ for the Canadian Arctic, due to the existence of specimens of difficult identification that might also be classified as D. cespitosa ssp. brevifolia. 7. Deschampsia cespitosa ssp. macrothyrsa (Tatew. et Ohwi) Tzvelev, Zlaki SSSR: 416. 1976. Deschampsia cespitosa var. macrothyrsa Tatew. et Ohwi, J. Jap. Bot. 17: 448. 1941. Deschampsia macrothyrsa (Tatew. et Ohwi) Kawano, Canad. J. Bot. 41: 740. 1963. type: Japan, Hab. Yezo, Onneto prope Nemuro, M.Tatewaki s .n. (SAPS, n.v.). Description: Plants 80–100 cm high; shoots thick with elongated rhizome-like lower internodes; ligules 4–5 mm long, membranaceous with margins hyaline; blades flat, 5–25 cm long, 2–3 mm wide, sparsely scabrous on adaxial side. Panicles open, lax, 25–35 cm long, branches scabrous; spikelets 4–5 mm long; upper glume slightly larger than the lower; awns curved, 2–3 mm long, inserted in or near the base of the lemma. Chromosome number: 2n = 26 (Probatova, 1984). Distribution: Russia: coast of the Sea of Okhotsk (near Magadan), Sakhalin, the Kuriles. Japan: Hokkaido. Ecology: Marshy sea coast and coastal meadows. Specimens examined:russia. Magadansky Region, in vicinity of Magadan city, Novaya Vesselaya settlement, meadow, M. Maksimova s.n., 27.vii.1970 (VLA); Sakhalinskaya Region, Kuriljsky district, Iturup Island, Burevestnik settlement, on the place of burned forest, E. Egorova et M. Aleksandrova s.n., 4.viii.1976 (MHA); Kurile Islands, Iturup Island, Burevestnik settlement, on the sea coast, V.N. Woroschilow 10897, 2.ix.1961 (MHA) Sakhalin Island, Nogliksky district, Nabilj Bay, on maritime sandy place, V.A. Nechajev, 28.viii.1977 (VLA); Sakhalin, Korsakovsky district, vicinity of Solovjevka settlement, marine terrasse, swampy meadow, N.S. Probatova 1723, 9.ix.1968 (VLA). japan. Hokkaido, prov. Tokachi, Bansei taiki-choo, coast and vicinity, sandy place, M. Furuse 6408, 18.vii.1974 (K). 8. Deschampsia cespitosa ssp. mezensis (Senjan.-Korcz. et Korcz.) Tzvelev, Fl. Europ. Chasti SSSR 1: 210. 1974. Deschampsia mezensis Senjan.-Korcz. et Korcz., Bot. Mater. Gerb. Bot. Institute Acad. Nauk SSSR 15: 31. 1953. type: Russia, Arkhangelsk district, Mezen river, A. Korchagin et M. Senyaninova-Korchagina, 2.vii.1949 (holotype LEU, n.v.). Description: Plants 70–80 cm high; lower internodes rhizome-like, elongated; sheaths glabrous with hyaline margins; ligules acute, 4–5 mm long, hyaline; blades 10–20 cm long, 2–3 mm wide, adaxially scabrous on top of nerves. Panicles open, 20–30 cm long, branches nearly glabrous with sparse prickles; spikelets 3–5 mm long, 2-flowered; lower glume lanceolate; upper glume broadly lanceolate; both glumes equal in size, 3–5 mm long, hyaline, membraneous only at the base and along the nerves; awns straight, 1.5–2.5 mm long, inserted at the base of the lemma. Chromosome number: 2n = 26 (Sokolovskaya & Probatova, 1975). Distribution: Russia: Arkhangelsk district, Komi Region, along Mezen and Vashka rivers. Ecology: Sandy unstable soils. Specimens examined:russia. Komi Republic, Mezen basin, sandbank near Vendenga, A. Korchagin et E. Ispolatov, 29.vii.1949 (LEU); Komi Republic, Intinsky district, Soviet Farm Gornyak, sandy bank of the Yunj-Yaga River, A.P. Sokolovskaya 77, 12.vii.1967 (VLA). 9. Deschampsia cespitosa ssp. obensis (Roshev.) Tzvelev, Fl. Evrop. Chasti SSSR 1: 210. 1974. Deschampsia obensis Roshev., Izv. Bot. Sada Akad. Nauk SSSR 30: 771. 1931. type: Russia, Obj River, Berezovskii post, Nakhodka Bay, A. Buschewich 39, vii. 1912 (holotype LE!). Description: Plants 40–60 cm high; lower internodes rhizome-like elongated, forming lax tussocks or also single shoots; ligules 3–5 mm long; blades flat or rather folded, 5–8 cm long, 0.5–1.5 mm wide. Panicles open, lax, 11–16 cm long, at least 1/3 or up to the half of the total length of the plant, with branches glabrous or sparsely scabrous, green; spikelets 5–7 mm long, 2–(3)-flowered, not numerous; glumes roughly equal, narrowly lanceolate (4)−5–7 mm long; awns straight to slightly bent and twisted, scabrous, 2.5–3.5 mm long, inserted between the middle and the lower part of the lemma and not exceeding the apex. Chromosome number: 2n = 42, 44, 52 (Zhukova & Petrovsky, 1975, 1980). Distribution: Russia. European and Siberian Arctic, lower parts and estuaries of the Obj and Yenisseisk Rivers. Ecology: Coastal sand dunes at sea level. Specimens examined:russia. Timanskaya tundra, A. Dedov, 4.viii.1928 (LE); Malozemelskaya tundra, A. Dedov, 18.viii.1930 (LE); Tobolsk prov. Berezow district, in arenosis ad fl. Polui prope Obdorsk, P. Pole 2710, 30.vi.1913 (LE, BM); in insulis Jenisseae fl., F. Schmidt 1866 (LE); Turukhansk district, Enisseisk, 71°53′, N.N. Kusnezow et W. W. Reverdatto (LE); Siberia, Yenisseisk, M. Brenner, 28.vii.1876 (LE); Tajmyr Peninsula, central part of peninsula, Byrranga Mountains, Fadjukuda River, sandy slope close to the river, I.N. Pospelov 97–275, 15.vii.1997 (MHA); Yakutia, delta of Lena River, sandy alluvium near river Erdeyalk, A. Tolmachow et al. 21.vii.1956 (LE); Siberia arctica, ad ostia Lenae, Dr Bunge s.n., viii.1884 (BM); Yakutia, Nizhne-Kolymski District, 10 km NW from Pokhodsk settlement, Galaktionova s.n., 2.viii.1980 (VLA); Chukotka Region, coast of the East Siberian Sea, Medvezhka River, tidal zone near the mouth of the river, partially flooded, V.V. Petrovsky s.n., 24.viii.1972 (VLA); Magadansky Region, Chaunsky District, Ustj-Chaun, field station Rossypnoi, M. Maximova 213–4000, 10.ix.1971 (VLA); Herb. Institute Bot. Academiae Sc. URSS 250, 19.vii.1964 (M). Observation 1: Ssp. obensis (Roshev.) Tzvelev and ssp. mezensis (Senjan.-Korcz. et Korcz.) Tzvelev have the lower internodes elongated rhizome-like, are morphologically similar, inhabit the same environment (sandbanks of rivers and coastal dunes) and have parapatric geographical distributions. It was not possible to study the type specimen of D. mezensis Senjan.-Korcz et Korcz. at LEU, apparently missing during renovation of the herbarium. Observation 2: Hybrids between D. cespitosa ssp. cespitosa and D. cespitosa ssp. obensis (Roshev.) Tzvelev have been found in Yakutia, in the lower part of Lena River (N.S. Probatova 3770, VLA) and in Magadansky Region, Ajan-Uriakh River, an tributary of Kolyma River (S.S. Kharkevich s.n., 13.vii.1989, VLA). 10. Deschampsia cespitosa ssp. orientalis Hultén, Kungl. Svenska Vetenskapsakad. Handl. 5: 109. October 1927. D. sukatschewii ssp. orientalis (Hultén) Tzvelev, Fl. Russian Arctic: 157. 1995. type: Russia, Kamchatka australis: Akhomten Bay, 275 m, E. Hultén 1362, 13.ix.1920 (holotype S! isotype LE!). Deschampsia cespitosa ssp. festucifolia Honda Bot. Mag. Tokyo 41: 635, 666. November 1927. type: Japan, Kiusiu, in Monte Kirishima, prov. Osumi, K. Mayebara 250, 1919 (SAPS?, n.v.). Deschampsia pumila (Griseb.) Ostenf. Meddel. Gronland 64: 169. 1927. type: Russia, Kamchatka (Isotype LE!). Deschampsia cespitosa var. microstachya Roshev., Izv. Glavn. Bot. Sada SSSR 28: 381. 1929. type: Russia, Siberia australis, in Transbaicalia frequens (holotype LE!). Deschampsia cespitosa var. paramushirensis (Honda) Tzvelev Grasses of the Soviet Union 1: 417. 1976. D. paramushirensis Honda J. Fac. Sci. University Tokyo, Sect. 3, Bot. 3 (1): 140. 1930. type: Russia, Kurile Islands, Kamogawa, ins. Paramushir, Y. Kudo 5272, 1920 (isotype TUS, n.v) non D. atropurpurea var. paramushirensis. Deschampsia cespitosa var. komarovii (A.V. Vassil.) Tzvelev Grasses of the Soviet Union 1: 417. 1976. D. komarovii A.V. Vassil. Bot. Mat. 8 (5): 68. 1940. type: Russia, Okhotsk, Ayan Region, valley of the Lantar river, A.V. Vassiliev 412, 26.viii.1935 (isotype LE!). Description: Plants 10–60 cm high; ligules oblong, 2–3 mm long; blades flat to folded, 8–20 cm long, 1–2 mm wide. Panicles lax, rarely slightly contracted, 6–18 cm long, with branches flexuose, scabrous; spikelets 3–5 mm long, 2–(3)-flowered; glumes slightly unequal, 3–5 mm long; awns straight to slightly undulated, 3–4 mm, inserted in or below the middle of the lemma, normally exceeding the glumes. Chromosome number: 2n = 26, 27–28, 52 (Sokolovskaya & Probatova, 1975; Osada, 1993; Takhtadzjan, 1993; as D. cespitosa ssp. festucifolia). Distribution: Russia. Kamchatka, Yakutia. Japan. Hokkaido, Honshu, Kiusiu. Ecology: Sandy coastal soils, flooded plains, tidal zones. Specimens examined:russia. Yakutiya, Tomponoskii, W. Iwanowa 64/3, 3.vii.1954 (LE); Yakutia, near village Nizhne, Rasputin, 4.vii.1903 (LE); Kamchatka, V. Komarov, 11.viii.1908 (LE); Kamchatka, P. Novograblenov 1088 and 1052, 23.vii.1929 (LE). japan. Pref. Iwate, prov. Rikuchyuu, Mt. Yakeishi-dake, 1400–1500 m, M. Furuse 34155, 24.vii.1958 (K); Pref. Nagano, prov. Shinano, Mt. Kiso-koma, lower part from Senjoo-jiki, M. Furuse 17177, 2.ix.1933 (K); Pref. Tochigi, prov. Shimotsuke, Sai-lake, from Senjyu-ga-hama side of Chyuusenji-lake, c. 1280 m, M. Furuse 42284, 2.vii.1964 (K). Observation 1:Hultén (1927: 109) described this taxon with contracted panicles, in spite of the fact that the holotype and isotype specimens (conserved in S and LE) show plants with open and lax panicles, while the only plants with contracted panicles are not completely developed. In the original description, Hultén regarded this taxon as restricted in its distribution to eastern Siberia. Later, in 1941, he considered that it represented an arctic-alpine type with a wider circumpolar distribution. Hultén (1941) also noted the existence of intermediate plants between ssp. orientalis and ssp. borealis (Traut.) Á. Löve et D. Löve, a taxon widely distributed in the Russian high arctic and which differs from ssp. orientalis by the smaller plants and more contracted panicles. Although there are intermediates between both types, they can be differentiated. The epithet orientalis is reserved here for taller plants with open panicles growing in eastern Siberia, Kamchatka and Japan, while borealis is restricted to the high arctic. Another related taxon native to Japan, D. cespitosa ssp. festucifolia Honda, with 2n = 27–28 (Osada, 1993), is morphologically very similar to orientalis and was synonymized by Osada (1993), as followed here. Observation 2: Tzvelev in 1995 made the combination D. sukatschewii ssp. orientalis (Hultén) Tzvelev. Observation 3: Both epithets festucifolia Honda and orientalis Hultén were published in 1927, although Hultén's Flora of Kamchatka was printed in October and the Botanical Magazine of Tokyo in November. 11. Deschampsia cespitosa ssp. parviflora (Thuill.). K. Richt., Pl. eur.1: 1890. Aira parviflora Thuill., Fl. env. Paris ed. 2 1: 38. 1799. Deschampsia cespitosa var. parviflora (Thuill.) Coss. et Germ., Obs. pl. env. Paris ed. 2: 806. 1861. type: France, Paris, hab. in sylvis, parc de Vincennes, de St. Maur, de côté de la Marne, fleurit in Juillet (holotype P?, n.v.). Aira altissima Lam. Fl. Franc. ed 3: 581. 1778. Deschampsia cespitosa var. altissima (Lam.) Volkart, Hegi Ill. Fl. Mitt-Eur. 1: 244. 1907. Description: Plants 80–140 cm tall; ligules 3–10 mm long; blades 10–34 cm long, 1.5–3 mm long. Panicles open and lax, 15–37 cm long; spikelets 2–3,5 mm long, 1–2-flowered; lower glume 2–3 mm long; upper glume 2–3.5 mm long, lemma 2–3 mm long; awn normally straight, 1–3 mm long, inserted in the basal third of the lemma. Chromosome number: 2n = 26, 28 (Kerguelén, 1993). Distribution. Russia. European part, Caucasus, west Siberia. Lithuania. Ukraine. Ecology: Damp mountain woods, shaded places, also in open wet meadows. Specimens examined:russia. Leningrad region, Boksitogorsky district, Alnus woods, A. Borodina 204, 11.viii.1971 (LEU); Podporoschsky district, Lake Onega, E. Tepljakova 125, 15.viii.1975 (LEU); Tosnensky district, Betula and Alnus mixed forest, A. Lavrenko, 17.vii.1975 (LEU); prov. Mosqua, districtus Zvenigorod prope Lusyno, piceetum humidum, V.V. Makarov 10, 2.vii.1969 (K); Moskovskaya Region, Serpukhovsky district, vicinity of Danki settlement, Prioksko-Terrasny Reserve, forest with Pinus and Quercus, S.S. Kharkevich et T.G. Buch, 10.vi.1973 (VLA); Siberia, Tomsk, M. Gandoger 1904 (BM). lithuania. Kaunassky district, Kaunassky reservoir near the village Verschuglisis, wet meadow in Alnus forest, V. Makarov s.n., 21.vi.1986 (MHA). ukraine. Kijevskaya Region, in vicinity of Irpenj town, Alnus forest, meadow, S.S. Kharkevich, 6.vii.1969 (VLA). Observation: Easily distinguished by its large pale green panicles and numerous small spikelets, often 1-flowered. 12.Deschampsia cespitosa ssp. submutica (Trautv.) Tzvelev Arct. Fl. SSSR 2: 84. 1964. Deschampsia cespitosa var. submutica (Trautv.) Tzvelev Zlaki SSSR: 282. 1976. Aira cespitosa var. submutica Trautv., Trudy Glavn. Bot. Sada 5: 141. 1877. D. sukatschewii ssp. submutica (Trautv.) Tzvel. Fl. Russ. Arctic: 156. 1995. type: Russia, Siberia, ad fl. Olenek, haud procul ab ostio fl. Maigda mediae, A. Czekanowski et F. Müller, 15.vii.1874 (holotype LE!). Description: Plants 35–45 cm high; sheaths glabrous with margins partially hyaline close to the ligule; ligules obtuse, 2–3 mm long; blades 2–5 cm long, 1–1.5 mm wide. Panicles rather contracted 8–11 cm, with branches curved, appressed, erect, glabrous or with a few sparse short hairs; spikelets 2–3 mm long, 2-flowered; lower glume lanceolate; upper glume broadly lanceolate; both glumes 2–3 mm long; lemmas 2–3 mm long, 4–5 toothed, teeth equal; awn straight, 1 mm long, weak, inserted in the middle of the lemma. Chromosome number: 2n = ? Distribution: Russia. Siberia, Olenek River. Ecology: Sandy riverside soils. Observation: Subspecies known only by the type specimen, distinguished by its very short awns and rather contracted panicles. 13. Deschampsia cespitosa ssp. sukatschewii (Popl.) Chiapella & Probatova comb. nov. Aira sukatschewii Popl. Ocherk. Fitosots. i Fitogeogr: 382. 1929. Deschampsia sukatschewii (Popl.) Roshev., Fl. URSS 2: 246. 1934. type: Russia, Transbaicalia inter fl. Nercza et fl. Kuenga, ad ripam fl. Olov, V.N. Sukatschew et G. Poplavskaya 1790, 13.viii.1911 (holotype LE!). Description: Plants 40–70 cm high; shoots erect forming tussocks without old sheaths at the base; ligules acute, 2–3 mm long; blades 9–15 cm long, conduplicate to flat, 1–2 mm wide, soft, green, glabrous to sparsely scabrous. Panicles rather open, 10–15 cm long, with numerous spikelets, from pale green to coloured, branches thin, long, erect to nodding, glabrous; spikelets 2.5–3.5 (4) mm long, 2-flowered; lower glume 2.5–3 mm long, narrowly lanceolate; upper glume 2.5–3.5 mm long, broadly lanceolate; lemma lanceolate with lateral nerves evident, teeth equal; awn straight to slightly bent, 2–3 mm, inserted in the basal third of the lemma. Chromosome number: 2n = 24, 26, 42, 42 + 1B, 52 (Takhtadzjan, 1993). Distribution: Russia: East Siberia, Yakutia region, southern part of the Russian Far East. Ecology: Sandy and gravelly shores of rivers, growing among bushes, wet meadows. Specimens examined:russia. Yakutia Region, Oljokminsky district, valley of Oljokma River, 10 km upstream the mouth of its affluent Chara, on gravel, M. Karavajev, 20.vii.1952 (MHA); Amurskaya Region, Skovorodinsky district, in vicinity of abandoned settlement Sverbeyevo, Amur River, 5 km up the confluence with Oldoi River, sandy slope on the riverside, A.E. Kozhevnikov s.n., 24.vi.1989 (VLA); Amurskaya Region, Zeyskij district, valley of Zeya River 12 km up Zeya town, stony slope in the valley of the river, C.D. Schlotgauer s.n., 25.vii.1971 (VLA); Magadanskaya Region, Khassynsky district, Talaja settlement, swampy Larix wood, A. Khokhrjakov, M.Mazurenko, 9.viii.1969 (MHA); Khabarovsky Territory, Tuguro-Chumikansky district, Nizhnjaja Maya River, Magan rivulet, gravelly shore, C.D. Sclotgauer 718, 25.vii.1970 (MHA). Observation: Ssp. sukatschewii seems to be a vicariant of ssp. cespitosa, especially in the Russian Far East where ssp. cespitosa is found as introduced in disturbed habitats (close to railways, near settlements, pasture lands, etc.). 14.Deschampsia cespitosa ssp. turczaninowii (Litv.) Tzvelev, Zlaki SSSR: 281. 1976. Aira turczaninowii Litv., Spisok Rast. Gerb. Russ. Fl. 8: 158. 1922. Deschampsia turczaninowii Litv. Spisok Rast. Gerb. Russ. Fl. 158, nom. altern. 1922. type: Russia, Baikal Lake, sandy-stony coast of Baikal Lake, on the Svyatoj Nos Peninsula, near Makarovka River, I. Larin et G. Kanevskii 2712, 30.vi.1916 (holotype LE! isotype S!). Description: Plants 20–60 cm high; ligules acute, 6–9 mm long; blades conduplicate, 4–8 cm long, 1–1.5 mm wide, with small prickles on top of nerves at the adaxial side. Panicles oblong, slightly contracted, 8–12 cm long, with branches erect, appressed, scabrous; spikelets 2–7 mm long, 2–(3)-flowered; lower glume 2–5 mm long; upper glume 3–7 mm long; awn straight, 5–8 mm long inserted in the lower third of the lemma and exceeding the glumes. Chromosome number: 2n = ? Distribution: Russia: Baikal Lake. Endemic. Ecology: Sandy rocky soils by the shore of Baikal Lake. Specimens examined:russia. Herb. Instituti Bot. Ac. Sc. U.S.S.R., 7.viii.1967 (K); Baikal lake, shore near mouth of river Tarkulik, N. Troyskaya, 18.vi.1988 (LE); Zabaikaliye province, Barguzinski district, shore of Baikal Lake, Peninsula Svyatoj Nos, between sand and stones, I. Larin et G. Kanevskij 308, 29.vi.1916 (LE); Irkutsk province, shore of Baikal Lake, Pestchanaya Bay, rare, I. Larin 60, 18.vi.1915 (LE). DISCUSSION In a study on Agrostis tenuis Sibth. in central Wales, Bradshaw (1959) found that despite a continuous distribution, distances of 50 m or less were enough to isolate populations. These populations showed environment-induced morphological differentiation but the differences were not considered worthy of recognition in a taxonomic scheme, due to the impossibility in delimiting non-arbitrary units (Bradshaw, 1959). In the present case the large Russian, nearly continuous, population of Deschampsia cespitosa (L.) P. Beauv., a species with a high morphological variability (Lawrence, 1945; Kawano, 1963, 1966; Ward, 1969; Pearcey & Ward, 1972; Chiapella, 2000a), inhabits a vast territory with high environmental heterogeneity, thus being subject to evolutionary pressure in several different directions. The existence of different morphological variants, however, due to tolerance plasticity (Morriset & Boutin, 1984: 295), and as long as they are related to a more or less restricted and defined geographical distribution, allows for a tentative division into subspecies as defined by Du Rietz (1930). The distribution pattern of the taxa included in the D. cespitosa complex in Russia resembles that of another, mostly Eurasian, complex with a world-wide distribution, Poa pratensis L. (Tzvelev, 1976). Several taxa of the P. pratensis complex were treated as subspecies by Tzvelev (1976) and have similar geographical distributions. A taxon with a widespread distribution is Poa pratensis ssp. pratensis, an extremely polymorphic species which is found in a wide range of environments in Eurasia and also North America, but is introduced in the Russian Far East and naturalized in Sakhalin, the Kuriles and Kamchatka. Other taxa are more or less confined, like Poa pratensis ssp. colpodea (Th. Fries) Tzvel., a pseudoviviparous taxon described from Spitzbergen which is more or less restricted to the high Arctic and which overlaps partially with D. cespitosa ssp. alpina, and like Poa pratensis ssp. sabulosa (Roshev.) Tzvel., a probable endemic to the Baikal lake and parts of nearby Mongolia, resembling the distribution of D. cespitosa ssp. turczaninowii. In the D. cespitosa complex, diploids and polyploids forms often coexist in sympatry, a characteristic that Stebbins (1975) has deemed as typical of young species complexes that still have not completed their development. The level of ploidy was not clearly related to the geographical distribution, except for a weak tendency of some polyploid forms (ssp. borealis, ssp. alpina and ssp. beringensis) to inhabit harsher habitats like the arctic, the high mountains and coastal marshes. Some of the taxa considered in this work showed several levels of ploidy (Table 1), suggesting the existence of highly adaptive phenotypes acting like a mask under which several different genomes would coexist (Barlow-Irick, 1997), thus avoiding diversification. The arctic taxa are adapted to their environment in a similar manner to the adaptation to high altitudes, which is characterized by a decrease in plant size and shorter leaves and inflorescences (Schmid, 1992). The existence of a latitudinal environmental gradient is confirmed by the height of the arctic taxa (ssp. borealis, ssp. brevifolia) which tend to be smaller than more southerly distributed taxa (ssp. macrothyrsa, ssp. mezensis, ssp. obensis; Table 1). Table 1. Comparative summary of the accepted taxa. Plant size, length of panicles and length of blades are given in cm. Size of spikelets, width of blades (measured unfolded) and length of ligules are given in mm. Sources for the chromosome numbers are given in the text Taxa . Plan height . Panicle length . Spikelet . Blades (length × width) . Ligule length . Awn . Pseudovivipary . Chromosome number (2n =) . ssp. alpina 8–40 5–15 2–5.5 2–5 × 0.5–2 2–6 1–3 yes 26, 35–38, 38–39, 39, 41, 48, 49, 49 + 2B, 49–52, 50, 52, 56 ssp. beringensis 25–80 9–20 4–7 5–12 × 2–4 8–12 3–6 no 26, 42 ssp. borealis 5–30 3–6 3–4 5–7 × 0.5–1.5 3–4 2–3 no 24–26, 26, 28, 42, 52 ssp. brevifolia 9–55 2.5–15 3–6 2–10 × 0.5–1.5 1.5–4 2–4 no 26, 27, 28, c. 50, 52 ssp. cespitosa 5–150 3–37 2–7 2–34 × 1–5 1.5–12 1–8 rare 18, 24, 24–25, 26, 26 + 1B, 27, 28, c. 39, 52 ssp. glauca 30–50 10–14 2–4.5 4–8.5 × 1–2 3–4 1.5–3.5 rare 26, 48–50, c. 52, 52 ssp. macrothyrsa 80–100 25–35 4–5 5–25 × 2–3 4–5 2–3 no 26 ssp. mezensis 70–80 20–30 3–5 10–20 × 2–3 3–4.5 1.5–2.5 no 26 ssp. obensis 40–60 11–16 5–7 5–8 × 0.5–1.5 3–5 2.5–3.5 no +42, +44, 52 ssp. orientalis 10–60 6–18 3–5 8–20 × 1–2 2–3 3–4 no 26, 27–28, 52 ssp. parviflora 80–140 15–37 2–3.5 10–34 × 1.5–3 3–10 1–3 no 26, 28 ssp. submutica 35–45 8–11 2–3 2–5 × 1–1.5 2–3 1–1.5 no unknown ssp. sukatschewii 40–70 10–15 2.5–3.5 9–15 × 1–2 2–3 2–3 no 24, 26, 42, 42 + 1B, 52 ssp. turczaninowii 20–60 8–12 2–7 4–8 × 1–1.5 6–9 5–8 no unknown Taxa . Plan height . Panicle length . Spikelet . Blades (length × width) . Ligule length . Awn . Pseudovivipary . Chromosome number (2n =) . ssp. alpina 8–40 5–15 2–5.5 2–5 × 0.5–2 2–6 1–3 yes 26, 35–38, 38–39, 39, 41, 48, 49, 49 + 2B, 49–52, 50, 52, 56 ssp. beringensis 25–80 9–20 4–7 5–12 × 2–4 8–12 3–6 no 26, 42 ssp. borealis 5–30 3–6 3–4 5–7 × 0.5–1.5 3–4 2–3 no 24–26, 26, 28, 42, 52 ssp. brevifolia 9–55 2.5–15 3–6 2–10 × 0.5–1.5 1.5–4 2–4 no 26, 27, 28, c. 50, 52 ssp. cespitosa 5–150 3–37 2–7 2–34 × 1–5 1.5–12 1–8 rare 18, 24, 24–25, 26, 26 + 1B, 27, 28, c. 39, 52 ssp. glauca 30–50 10–14 2–4.5 4–8.5 × 1–2 3–4 1.5–3.5 rare 26, 48–50, c. 52, 52 ssp. macrothyrsa 80–100 25–35 4–5 5–25 × 2–3 4–5 2–3 no 26 ssp. mezensis 70–80 20–30 3–5 10–20 × 2–3 3–4.5 1.5–2.5 no 26 ssp. obensis 40–60 11–16 5–7 5–8 × 0.5–1.5 3–5 2.5–3.5 no +42, +44, 52 ssp. orientalis 10–60 6–18 3–5 8–20 × 1–2 2–3 3–4 no 26, 27–28, 52 ssp. parviflora 80–140 15–37 2–3.5 10–34 × 1.5–3 3–10 1–3 no 26, 28 ssp. submutica 35–45 8–11 2–3 2–5 × 1–1.5 2–3 1–1.5 no unknown ssp. sukatschewii 40–70 10–15 2.5–3.5 9–15 × 1–2 2–3 2–3 no 24, 26, 42, 42 + 1B, 52 ssp. turczaninowii 20–60 8–12 2–7 4–8 × 1–1.5 6–9 5–8 no unknown Open in new tab Table 1. Comparative summary of the accepted taxa. Plant size, length of panicles and length of blades are given in cm. Size of spikelets, width of blades (measured unfolded) and length of ligules are given in mm. Sources for the chromosome numbers are given in the text Taxa . Plan height . Panicle length . Spikelet . Blades (length × width) . Ligule length . Awn . Pseudovivipary . Chromosome number (2n =) . ssp. alpina 8–40 5–15 2–5.5 2–5 × 0.5–2 2–6 1–3 yes 26, 35–38, 38–39, 39, 41, 48, 49, 49 + 2B, 49–52, 50, 52, 56 ssp. beringensis 25–80 9–20 4–7 5–12 × 2–4 8–12 3–6 no 26, 42 ssp. borealis 5–30 3–6 3–4 5–7 × 0.5–1.5 3–4 2–3 no 24–26, 26, 28, 42, 52 ssp. brevifolia 9–55 2.5–15 3–6 2–10 × 0.5–1.5 1.5–4 2–4 no 26, 27, 28, c. 50, 52 ssp. cespitosa 5–150 3–37 2–7 2–34 × 1–5 1.5–12 1–8 rare 18, 24, 24–25, 26, 26 + 1B, 27, 28, c. 39, 52 ssp. glauca 30–50 10–14 2–4.5 4–8.5 × 1–2 3–4 1.5–3.5 rare 26, 48–50, c. 52, 52 ssp. macrothyrsa 80–100 25–35 4–5 5–25 × 2–3 4–5 2–3 no 26 ssp. mezensis 70–80 20–30 3–5 10–20 × 2–3 3–4.5 1.5–2.5 no 26 ssp. obensis 40–60 11–16 5–7 5–8 × 0.5–1.5 3–5 2.5–3.5 no +42, +44, 52 ssp. orientalis 10–60 6–18 3–5 8–20 × 1–2 2–3 3–4 no 26, 27–28, 52 ssp. parviflora 80–140 15–37 2–3.5 10–34 × 1.5–3 3–10 1–3 no 26, 28 ssp. submutica 35–45 8–11 2–3 2–5 × 1–1.5 2–3 1–1.5 no unknown ssp. sukatschewii 40–70 10–15 2.5–3.5 9–15 × 1–2 2–3 2–3 no 24, 26, 42, 42 + 1B, 52 ssp. turczaninowii 20–60 8–12 2–7 4–8 × 1–1.5 6–9 5–8 no unknown Taxa . Plan height . Panicle length . Spikelet . Blades (length × width) . Ligule length . Awn . Pseudovivipary . Chromosome number (2n =) . ssp. alpina 8–40 5–15 2–5.5 2–5 × 0.5–2 2–6 1–3 yes 26, 35–38, 38–39, 39, 41, 48, 49, 49 + 2B, 49–52, 50, 52, 56 ssp. beringensis 25–80 9–20 4–7 5–12 × 2–4 8–12 3–6 no 26, 42 ssp. borealis 5–30 3–6 3–4 5–7 × 0.5–1.5 3–4 2–3 no 24–26, 26, 28, 42, 52 ssp. brevifolia 9–55 2.5–15 3–6 2–10 × 0.5–1.5 1.5–4 2–4 no 26, 27, 28, c. 50, 52 ssp. cespitosa 5–150 3–37 2–7 2–34 × 1–5 1.5–12 1–8 rare 18, 24, 24–25, 26, 26 + 1B, 27, 28, c. 39, 52 ssp. glauca 30–50 10–14 2–4.5 4–8.5 × 1–2 3–4 1.5–3.5 rare 26, 48–50, c. 52, 52 ssp. macrothyrsa 80–100 25–35 4–5 5–25 × 2–3 4–5 2–3 no 26 ssp. mezensis 70–80 20–30 3–5 10–20 × 2–3 3–4.5 1.5–2.5 no 26 ssp. obensis 40–60 11–16 5–7 5–8 × 0.5–1.5 3–5 2.5–3.5 no +42, +44, 52 ssp. orientalis 10–60 6–18 3–5 8–20 × 1–2 2–3 3–4 no 26, 27–28, 52 ssp. parviflora 80–140 15–37 2–3.5 10–34 × 1.5–3 3–10 1–3 no 26, 28 ssp. submutica 35–45 8–11 2–3 2–5 × 1–1.5 2–3 1–1.5 no unknown ssp. sukatschewii 40–70 10–15 2.5–3.5 9–15 × 1–2 2–3 2–3 no 24, 26, 42, 42 + 1B, 52 ssp. turczaninowii 20–60 8–12 2–7 4–8 × 1–1.5 6–9 5–8 no unknown Open in new tab CONCLUSIONS Due to extensive overlapping in morphological diagnostic characters with D. cespitosa, it is suggested that D. alpina (L.) Roem. et Schult., D. borealis (Trautv.) Roshev., D. brevifolia R. Br., D. glauca C. Hartm., D. macrothyrsa Tatew. et Ohwi, D. obensis Roshev. should be considered as subspecies of D. cespitosa. All them were considered species by Tzvelev (1964) and Czerepanov (1995). Deschampsia vodopjanoviae O.D. Nikif. is synonymised with D. cespitosa (L.) P. Beauv. The present survey suggests that morphological characters alone do not provide a clear differentiation among the considered taxa, and lends support for a broad consideration of D. cespitosa and the approach followed by Tzvelev in 1974 and 1976, i.e. assigning subspecies level to the taxa with a more or less defined geographical distribution. It is, however, clear that in order to clarify definitely the relationships among the studied taxa and its placement in a taxonomic scheme an exhaustive molecular-based study would be necessary, particularly in the north-west Pacific area. Stebbins (1975) pointed to eastern Siberia as a probable centre of origin and radiation for species of the Elymus–Hordeum group, a hypothesis that the diversity of Deschampsia seems to support too. Phylogenetic relationships in Deschampsia and the Aveneae have not yet been fully clarified (Soreng & Davis, 1998, 2000), and the taxonomy of the genus remains in the making. ACKNOWLEDGEMENTS JC thanks the financial asistance received from the Royal Botanic Gardens, Kew, through the Latin American Fellowship (financed by the Mellon foundation), which allowed visits to B, BM, K, LE, LEU, S, and W. Alfred O'Passo (Bariloche) and Heimo Rainer (Vienna) helped with the preparation of the distribution map. Dmitry Geltman, Vyacheslav Byalt, Alexander Sennikov and Ivan Schanzer helped with the translation from Russian of numerous herbarium labels at LE and MHA. This work was completed with a fellowship of the ‘Consejo Nacional de Investigaciones Científicas y Técnicas’ (CONICET) from Argentina. 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GROLLE, RICLEF; SO, MAY LING

doi: 10.1046/j.1095-8339.2003.00176.xpmid: N/A

AbstractRiccia fruticulosa O.F.Müll., 1782 from Norway is a valid name, referring to Riccardia palmata (Hedw.) Carruth. In 1785 Dickson misidentified British plants of a blue Metzgeria as R. fruticulosa. The European blue species of Metzgeria is conspecific with M. violacea (Ach.) Dumort., which replaces M. fruticulosa auct. The true origin of the type of Jungermannia violacea Ach., 1805 is probably Tierra del Fuego (rather than Dusky Bay, New Zealand), where the species is widespread. Reports from Australasia, Asia and Africa are all erroneous. The blue colour of Jungermanniales is found only in living plants and is derived from the oil-bodies. In contrast, that of Metzgeria appears only after death; its biological function is unknown.

ROUX, J. P.

doi: 10.1046/j.1095-8339.2003.00146.xpmid: N/A

AbstractDryopteris katangaensis, a new species endemic to the Katango–Zambian centre of endemism (Congo-Zambezi watershed) in Zambia and the Democratic Republic of Congo is described and illustrated. The morphological differences between the new species and D. schimperiana are discussed, and the ferns endemic to the Katango–Zambian Centre are reviewed.